Acta Botanica Mexicana (2001), 54: 29-37

HABITAT AND RANGE EXTENSION OF COBAEA LUTEA (POLEMONIACEAE)

IN WESTERN MEXICO

EDUARDO SAHAGN-GODNEZ
JOS AQUILEO LOMEL-SENCIN

Jardn Botnico y Herbario

Universidad Autnoma de Guadalajara
Apartado postal 1-440
44100 Guadalajara, Jalisco, Mxico
esahagun@uagunix.gdl.uag.mx jalomeli@icb.gdl.uag.mx

AND

L. ALAN PRATHER

Department of Botany and Plant Pathology

Michigan State University
East Lansing, MI 48824-1312, USA
alan@msu.edu

ABSTRACT

For the first time we present evidence of the presence of Cobaea lutea D. Don in tropical
deciduous forest; we propose an hypothesis to explain this event and discuss the variety of habitats
occupied by this and other members of the Polemoniaceae. We analyze the extension of the geographic
limits of C. lutea ca. 350 km northwestwards from the Sierra Madre del Sur in Guerrero to the Eje
Neovolcnico in Jalisco. We also document the presence of densely long-villous pedicels as a
morphological variant of C. lutea, and present a description and an illustration.

Key words: Cobaea, Polemoniaceae, tropical deciduous forest, phenotypic plasticity, western
Mexico.

RESUMEN

Por primera vez se registra la presencia de Cobaea lutea D. Don en bosque tropical caducifolio;
se propone una hiptesis para explicar este evento y se discute la diversidad de hbitats ocupados
por ste y otros miembros de las Polemoniaceae. Se analiza la extensin de los lmites geogrficos
de C. lutea ca. 350 km hacia el noroeste, desde la Sierra Madre del Sur en Guerrero hasta el Eje
Neovolcnico en Jalisco. Se documenta la presencia de pedicelos densamente vilosos como una
variante morfolgica de C. lutea y se describe e ilustra este taxon.

Palabras clave: Cobaea, Polemoniaceae, bosque tropical caducifolio, plasticidad fenotpica,

occidente de Mxico.

29
Acta Botanica Mexicana (2001), 54: 29-37

INTRODUCTION

There are 18 known species in the genus Cobaea Cav. Of these, 10 occur in Mexico
and four, all in section Cobaea, are endemic to this country (Prather, 1999). The genus is
characterized by its scandent habit and pinnately compound leaves with the terminal leaflets
modified into tendrils. The flowers are large and campanulate, the pollen grains are reticulate,
and the fruits are septicidal (Prather, 1996).
The species of Cobaea are known to occur in humid montane forests of tropical
America (Standley, 1914; Gibson, 1970; Ippolito & Surez, 1998). The habitat extension of
C. lutea D. Don presented here is significant because it could indicate that the lack of
specimens in herbaria of this and perhaps other species of Cobaea from low elevation tropical
regions may be due in part to the oversight of collectors of these green-flowered plants in
the vine-entangled vegetation of tropical deciduous forests.
For many years Cobaea lutea was regarded as a species of South and Central
America, with its northernmost known record in Guerrero. Only recently have new populations
been found as far north as Zacoalco in Jalisco, ca. 350 km northwest of the Guerrero
populations. This distance is more significant than it would seem at first glance due to the
ruggedness of the terrain.
The objectives of this article are to 1) document the presence of Cobaea lutea in
tropical deciduous forest, 2) compare this to the habitats occupied by other polemoniaceous
genera, 3) propose an hypothesis to account for the habitat expansion, and 4) report a
significant extension in the geographic range to the northwest.

METHODS

In March 1997 fruits from a population of Cobaea located at Las Moras in the municipio
of Zacoalco, Jalisco were collected. Seeds were sown at Jorge Victor Eller Botanical Garden
that summer. Herbarium specimens were prepared when the plants flowered in October. The
following year, additional seeds from the initial collection were again sown. Photographs and
a line drawing were prepared from these cultivated plants. Additional herbarium material was
prepared from plants growing in the wild at Las Moras during a collection trip in October
1998. This material was used to adapt the description of C. lutea given in Prather (1999).
All herbarium specimens were deposited at Carlos Luis Daz Luna Herbarium of
Universidad Autnoma de Guadalajara (GUADA). Additional specimens of C. lutea housed
in other herbaria (CAS, F, GH, IBUG, K, MEXU, MO, MSC, NY, RSA, TEX, UC, US, WIS)
were also examined. Other herbaria were consulted but they had no specimens of C. lutea
for locations north of Guerrero (CHAPA, ENCB, UAMIZ and XAL).

RESULTS

A population of C. lutea at Las Moras was found mixed with other vines (Cucurbita
sp., Sechiopsis triquetra and Quamoclit sp.) in one of the main ravines stemming down from

30
 Sahagn-Godnez et al.: Habitat and Range Extension of Cobaea lutea in Western Mxico

the oak and pine forest areas located higher in the Sierra de Tapalpa. The plants were
abundant, although localized, in tropical deciduous forest with Lysiloma acapulcense,
Montanoa sp., Bursera spp., Ipomoea murucoides, Euphorbia cotinifolia, Ptelea trifoliata,
Viguiera quinqueradiata, Tillandsia dasyliriifolia and Tillandsia spp.
In an effort to explain the presence of Cobaea lutea in the drier habitats of western
Mexico, we propose the following hypothesis. Cobaea lutea in this region is likely a relict
from widespread ancestral populations that lived in a more humid climate and now is found
only in favored sites in ravines. We also believe that a combination of adaptations such as
an annual habit and a vigorous scandent form have allowed C. lutea to survive in a seasonally
drier environment. We present support for this hypothesis in the discussion.
The population located at Las Moras is currently the northernmost known. A
description of Cobaea lutea follows to aid in its identification. Range values in brackets were
measured from the specimens collected at Las Moras.

Cobaea lutea D. Don, Edinburg Phil. J. 10: 112. 1824. Fig. 1.

Annual vine, 5-8 m long. Stems usually slender, sometimes puberulent at the nodes.
Leaves alternate, pinnately compound, with 6 opposite or subopposite leaflets, the terminal
leaflet modified into a branched tendril, each branch terminated with two claws; rachis 23-
70 mm long; petiolules [2] 3-7 mm long; leaflets 24-110 [130] mm long, 11-60 mm wide,
elliptic to elliptic-oblanceolate, rarely obovate, glabrous or puberulent along the veins, margins
minutely ciliate and sometimes long-ciliate at the base, base acute to truncate to cordate,
apex acute to acuminate. Inflorescence a dichasial cyme of 1-5 flowers, subtended by
foliaceous bracts. Peduncles (2.3) 6.3-10.0 (15.0) cm long, glabrous or puberulent at the
base and apex. Pedicels (8.2) 14.3-25.0 cm long, glabrous or puberulent at the base and
apex, sometimes long-villous, coiled in fruit. Calyx segments 5, 19-28 mm long, 3.5-8.0 mm
wide, essentially distinct, green, rarely suffused with red or purple, lanceolate to lanceolate-
ovate, apex acute to acuminate, minutely ciliate and sometimes also long-ciliate. Corollas
green, rarely greenish-yellow or suffused with purple or red, puberulent or sometimes villous
externally on the upper tube and the base of the lobes, glabrous internally except for the
long-villous annulus; tube 17-37 mm long, 19-35 mm wide, campanulate; lobes 5, 13-30 mm
long, 10-18 mm wide, broadly ovate, narrowing to an acuminate to long-acuminate apex,
slightly imbricate. Stamens 5; filaments 34-80 mm long, adnate basally to the corolla tube
for 4-9 mm, emergent or slightly reflexed; anthers [6] 7-14 mm long, [1] 1.5-2.5 mm wide,
yellow, linear to lanceolate, versatile. Ovary superior, inserted on a large five-lobed nectary
disk, tricarpellate, trilocular, ovate, glabrous, septa thickened and fused into a central column
which is triangular in cross section; styles 65-78 (90) mm long; style branches 3, 8-13.5 mm
long, terete, papillae short, covering nearly the entire surface. Fruit a capsule, exceeding
the calyx, 20-58 mm long, septicidal, elliptic to narrowly elliptic. Seeds 6-21 per fruit, flat,
ovate to broadly ovate, 16-21 [24] mm long including the wing, [10] 11-12 [13] mm wide
including the wing; wings 2-4 mm wide.

Phenology. Flowering in September-December. Fruiting in October-March.

Distribution. Peru, El Salvador, Honduras, Guatemala and Mexico (Chiapas, Guerrero,

Colima, and Jalisco). This species occurs in cloud forests, semi-deciduous forests,

31
Acta Botanica Mexicana (2001), 54: 29-37

Fig. 1. Cobaea lutea. A. Portion of plant with inflorescence. B. Leaf with detached leaflets. C. Dissected
corolla showing insertion of stamens, long-villous annulus and nectary disk. D. Immature fruit with coiled
pedicel showing densely long-villous pubescence. E. Dehisced fruit showing persistent calyx, septal
column and filamentous placentae. F. Winged seed. All drawings from a plant cultivated at Jorge Victor
Eller Botanic Garden, except for B, from J. A. Lomel 2928.

32
 Sahagn-Godnez et al.: Habitat and Range Extension of Cobaea lutea in Western Mxico

tropical deciduous forest and forested foothills of the Pacific coastal regions of North and
South America; 300-2200 m above sea level.
Specimens seen: COLIMA: Rancho El Jabal 20 km (airline distance) N of Colima
in the SW foothills of the Volcn de Colima. From the Jabal main gate to Hacienda San
Antonio along the road to Comala. Near 19026.09'N 103043'W, A. C. Sanders 11946 w/L.
Vzquez V. (CAS, MEXU). JALISCO: Sierra de Manantln Biosphere Reserve, near Colima
boundary, zone of permanently cultivated areas intercalated with extensive chaparral, right
below SE edge of Cerro Toxn in bottom of arroyo Pitahayas, 1 km NE of Toxn, barely
N of Puerto de Toxn-La Loma crossroads, along El Sauz-San Pedro Toxn Rd. 13 km (airline
distance) NNE of Minatitln, 39 km due NW of Colima, T. Cochrane et al. 12268 (MSC,
WIS-2); Mpio. Zapotitln, rancho El Jabal, 22 km (airline) N of Colima in the SW foothills
of the Volcn de Colima, La Joya area, N of the airstrip and S of Arroyo Santa Cruz, E
of Cerro El Campanario, A. C. Sanders 11666 (CAS); Mpio. de Zacoalco, caada en ladera
al NW de Las Moras, bosque tropical caducifolio, J. A. Lomel-Sencin s. n. Octubre 1997
(GUADA); Mpio. de Zacoalco, Las Moras, caada ca. 200 m ro arriba desde la Capilla de
la ranchera (20009'30"N 103035'35"W), bosque tropical caducifolio, J. A. Lomel-Sencin
2928 (GUADA, IBUG, MEXU); ibid., J. A. Lomel-Sencin 3446 (GUADA). GUERRERO:
District of Montes de Oca, San Antonio, G. B. Hinton 11671 (F, GH, K, MO, NY, RSA, UC,
US); along Route 134 between Ciudad Altamirano and the intersection with Route 200,
19.2 km N of San Antonio, A. Prather 1220 with J. Soule (MEXU, TEX).

DISCUSSION

Most researchers agree that Cobaea is a genus of humid montane forests (Standley,
1914; Gibson, 1970; Ippolito & Surez, 1998), thus the new record from tropical deciduous
forest is notable. Prather (1999) lists the habitat of most Cobaea species as mesic, tropical
montane forests (often cloud forests), but notes that a few occur at lower altitudes. All Cobaea
species were reported from mesic habitats, but Prather collected C. rotundiflora at one site
in tropical deciduous forest in Guatemala at 975 m alt. (Prather 960, TEX). Cobaea
rotundiflora is found more typically in mesic environments at higher elevations in Chiapas
(Prather, 1999).
Likewise, C. lutea populations were known to occur only in more mesic habitats, from
cloud forests to mesic forests of the foothills of the Pacific Coastal region in Guatemala and
El Salvador. Thus, C. lutea and C. rotundiflora share a similar breadth of habitat, have
common centers of distribution (even though the extent of C. lutea distribution is much
broader), are morphologically similar, and are closely related based on molecular data
(Prather, 1999). Nonetheless, they are easily distinguished by calyx width, stamen length
relative to corolla length, corolla shape, anther position and length of style branches (Prather,
1996).
It is possible that C. lutea may be widespread in tropical deciduous forests but it had
not been collected due to the difficulty of locating the green-flowered plants in the dense
vegetation. It is remarkable that, at Las Moras, it was difficult for us to find the flowers, even
when we knew they were in front of us since we had located the population in flower when
we were on our way down, on the other side of the creek. Cobaea lutea is much easier

33
Acta Botanica Mexicana (2001), 54: 29-37

to detect when it is in fruit. The large capsules hanging from coiled pedicels evidence both
location and identity.
The species of Cobaea are generally more mesic than most other species of the
Polemoniaceae, including the other members of the subfamily Cobaeoideae. According to
molecular data from the chloroplast genome, the genus Bonplandia is the sister group to
the genus Cobaea (e.g. Johnson et al., 1996; Prather et al., 2000). Bonplandia is a monotypic
genus that occurs mainly in tropical deciduous forests (Rzedowski & Caldern de Rzedowski,
1995) from Tamaulipas and southern Sonora, south to Guatemala. The western South
American genera Cantua and Huthia are also known to be related to Cobaea (Grant, 1998;
Johnson et al., 1996; Prather et al., 2000) and occur mainly in dry shrublands and forests
of the Andes (Weberbauer, 1945; Brako & Zarucchi, 1993). Loeselia and Acanthogilia, are
less closely related to Cobaea, but are distributed primarily (Loeselia) or exclusively
(Acanthogilia) in Mexico. The species of Loeselia are found in a variety of habitats, from
dry forests, to deciduous forests, and deserts (Rzedowski & Caldern de Rzedowski, 1995)
and Acanthogilia is found in deserts of the Baja Peninsula (Day & Moran, 1986). The
remainder of the species of the family occur in a variety of habitats, from deserts to vernal
pools or stream banks, but the majority of species of the family are desert herbs of western
North America (Grant, 1998).
In Mexico, Cobaea lutea is currently known to occur in Chiapas, Guerrero, Colima
and Jalisco. It is expected in Oaxaca and Michoacn, which lie between known populations,
but as far as we know, no specimens or published reports exist to confirm its presence there.
Even where reported, C. lutea seems to be rare or at least little collected.
In Guerrero, the recently-compiled checklists that we reviewed do not include any
species of Cobaea (Fonseca & Lozada, 1993; Diego & Lozada, 1994; Lozada, 1994; Gual,
1995; Peralta, 1995; Verduzco & Rodrguez, 1995; Gallardo, 1996 and Vargas & Prez,
1996).
From the valley of Mexico, Caldern de Rzedowski (1985) had no record of the genus,
but Argelles et al. (1991) reported Cobaea scandens from nearby Quertaro. From the Bajo
region, a thoroughly explored area which includes Guanajuato, Quertaro and portions of
Michoacn, Rzedowski and Caldern de Rzedowski (1995) reported Cobaea scandens and
C. stipularis only. Labat (1995) did not list any species of Cobaea in Michoacn, and neither
did Barrios-Rodrguez and Medina-Cota (1996) from Sierra de Pachuca, in Hidalgo.
From Jalisco, no species of Cobaea were reported in the publications reviewed
(Rzedowski & McVaugh, 1966; Lott, 1985; Vzquez et al., 1995; Guerrero-Nuo & Lpez-
Coronado, 1997), except for Prather (1999).
As far as we know, the northernmost record of this species known in the literature
is a specimen collected by Cochrane et al. (No. 12268) in the Manantln Biosphere Reserve
near the limits with the state of Colima, reported by Prather (1999). However, he did not
discuss the significant range extension for this species, ca. 300 km northwest of the Guerrero
populations (Fig. 2). The specimens collected by Lomel (No. 2928, 3446) reported here
further extend the range northwestward, ca. 350 km from the Guerrero collections. This
population also occurs farther inland than other collections known from Mexico. The Jalisco-
Colima specimens are particularly important because they were collected in the Eje
Neovolcnico, a physiographic region different from the Sierra Madre del Sur, where the
Guerrero collections came from.

34
 Sahagn-Godnez et al.: Habitat and Range Extension of Cobaea lutea in Western Mxico

Fig. 2. Known collection sites of Cobaea lutea in Mexico. Chiapas (Sierra Madre de Chiapas);
Guerrero (Sierra Madre del Sur); Colima and Jalisco (Eje Neovolcnico).

We believe it is possible for C. lutea to occur farther north on the Pacific coast since
other species of Cobaea have been found as far north as Nuevo Len on the Atlantic slope
(Hinton & Hinton, 1995). More fieldwork is necessary to increase the presence of Cobaea
specimens in herbaria.
Morphologically, the specimens collected at Las Moras in Jalisco (Lomel No. 2928,
3446) and at Rancho El Jabal in Colima (Sanders No. 11946) are unusual. The plants show
densely long-villous pedicels, a condition previously unknown in C. lutea. Prather (1996)
reported hairs longer than 4 mm on the pedicels of C. lutea. However, those hairs were
never dense on the specimens examined. Whether the presence of densely long-villous
pedicels is a result of phenotypic plasticity of the plants growing in drier environments (i.e.,
in tropical deciduous forest) or to other causes, is a subject for future study.
In support of the hypothesis proposed above to explain the presence of Cobaea lutea
in tropical deciduous forest, we may say that this species appears to be sensitive to
microclimate because at Las Moras, the majority of the population is restricted to the ravine.
The few plants observed outside were precocious and started fruiting earlier. These plants
also showed a reddish coloration and looked depauperate. Furthermore, Cobaea lutea is
clearly an annual species. This habit allows it to bypass the severe shortage of water of
the tropical lowlands during the dry season.
Cobaea might have originated in central Mexico (Grant, 1959) at a time when the
climate in Mexico was cooler and wetter, as it was during much of the Cenozoic, when
temperate forest elements migrated south into Mexico and covered extensive areas (Graham,
1973). Cobaea probably diversified in this wide-ranging temperate forest.

35
Acta Botanica Mexicana (2001), 54: 29-37

When the climate in northern Latin America became warmer and drier in recent times,
the populations of Cobaea receded together with the temperate forests to their present
location, and are now present on the Pacific slope in western Mexico only as relicts in favored
areas.
The presence of a few populations of Cobaea in tropical deciduous forest represents
some of these relicts that have been able to survive in moist creeks despite the heat, the
severe dry season of up to eight months, and the presence of other aggressive vines.
In this respect, we think that the annual, scandent habit, winged seeds and rapid
vigorous growth have played a role in the competitive ability and ultimately the survivorship
of Cobaea lutea in tropical habitats.

ACKNOWLEDGMENTS

We are thankful to Ren Len-Maldonado for his valuable help in the course of this
work. We would also like to thank Jorge Flores at Facultad de Ciencias Naturales y
Agropecuarias, Universidad Autnoma de Guadalajara, for fieldwork support. We are
indebted to herbarium curators who kindly provided information and access to their
specimens and to two anonymous reviewers, whose valuable suggestions are here gratefully
acknowledged.

LITERATURE CITED

Argelles, E., R. Fernndez & S. Zamudio. 1991. Listado florstico preliminar del estado de Quertaro.
Flora del Bajo y de Regiones Adyacentes II: 1-155. Instituto de Ecologa, A.C. Ptzcuaro,
Michoacn.
Barrios-Rodrguez, M. A. & J. M. Medina-Cota. 1996. Estudio florstico de la Sierra de Pachuca, estado
de Hidalgo. Instituto Politcnico Nacional. Mxico, D.F. 140 pp.
Brako, L. & J. L. Zarucchi. 1993. Catalogue of the flowering plants and Gymnosperms of Peru. Monogr.
Syst. Bot. Missouri Bot. Gard. Vol. 45. St. Louis. 1286 pp.
Caldern de Rzedowski, G. 1985. Polemoniaceae. In: Rzedowski & Rzedowski (eds.) Flora fanerogmica
del Valle de Mxico. Vol. II. Escuela Nacional de Ciencias Biolgicas and Instituto de Ecologa.
Mxico, D.F. pp. 257-262.
Day, A. G. & R. Moran. 1986. Acanthogilia, a new genus of Polemoniaceae from Baja California, Mexico.
Proc. California Acad. Sci. 44: 111-126.
Diego, N. & L. Lozada. 1994. Estudios florsticos en Guerrero. No. 3. Laguna de Tres Palos. Facultad
de Ciencias, Universidad Nacional Autnoma de Mxico. Mxico, D.F. 29 pp.
Fonseca, R. M. & L. Lozada. 1993. Estudios florsticos en Guerrero. No. 1. Laguna de Coyuca. Facultad
de Ciencias, Universidad Nacional Autnoma de Mxico. Mxico, D.F. 23 pp.
Gallardo, C. 1996. Estudios florsticos en Guerrero. No. 8. Parque ecolgico La Vainilla. Zihuatanejo,
Guerrero. Facultad de Ciencias, Universidad Nacional Autnoma de Mxico. Mxico, D.F. 61
pp.
Gibson, D. N. 1970. Polemoniaceae. In: Standley, P. C. & L. O. Williams (eds.) Flora of Guatemala.
Fieldiana, Bot. 24(9) parts I and II: 85-93.
Graham, A. 1973. History of the arborescent temperate element in the northern Latin American biota.
In: Graham, A. (ed.). Vegetation and vegetational history of northern Latin America. Elsevier
Scientific Publishing Company. Amsterdam. pp. 301-314.

36
 Sahagn-Godnez et al.: Habitat and Range Extension of Cobaea lutea in Western Mxico

Grant, V. 1959. Natural history of the Phlox family: Systematic Botany. The Hague: Martinus Nijhoff.
280 pp.
Grant, V. 1998. Primary classification and phylogeny of the Polemoniaceae, with comments on molecular
cladistics. Amer. J. Bot. 85: 741-752.
Gual, M. 1995. Estudios florsticos en Guerrero. No. 6. El Can del Zopilote. rea Venta Vieja. Facultad
de Ciencias, Universidad Nacional Autnoma de Mxico. Mxico, D.F. 39 pp.
Guerrero-Nuo, J. J. & G. A. Lpez-Coronado. 1997. La vegetacin y la flora de la Sierra de Quila,
Jalisco, Mxico. Universidad de Guadalajara. Guadalajara, Jalisco. 135 pp.
Hinton, J. & G. S. Hinton 1995. Checklist of Hintons collections of the flora of south central Nuevo
Len and adjacent Coahuila. Acta Bot. Mex. 30: 41-112.
Ippolito, A. & A. V. Surez. 1998. Flowering phenology and pollination of Cobaea aschersoniana
(Polemoniaceae). Biotropica 30(1): 145-148.
Johnson, L. A., J. L. Schultz, D. E. Soltis, & P. S. Soltis. 1996. Monophyly and generic relationships
of Polemoniaceae based on matK sequences. Amer. J. Bot. 83: 1207-1224.
Labat, J. N. 1995. Vgtation du nord-ouest du Michoacn, Mexique. Flora del Bajo y de Regiones
Adyacentes VIII. Instituto de Ecologa, A.C. Ptzcuaro, Michoacn. 401 pp.
Lott, E. J. 1985. Listados florsticos de Mxico. III. La Estacin de Biologa Chamela, Jalisco. Herbario
Nacional. Instituto de Biologa, Universidad Nacional Autnoma de Mxico. Mxico, D.F. 30
pp.
Lozada, L. 1994. Estudios florsticos en Guerrero. No. 2. Laguna de Mitla. Facultad de Ciencias,
Universidad Nacional Autnoma de Mxico. Mxico, D.F. 44 pp.
Peralta, S. 1995. Estudios florsticos en Guerrero. No. 5. Can del Zopilote (rea Papalotepec).
Facultad de Ciencias, Universidad Nacional Autnoma de Mxico. Mxico, D.F. 37 pp.
Prather, L. A. 1996. Three new species of Cobaea (Polemoniaceae). Brittonia 48(1): 111-119.
Prather, L. A. 1999. Systematics of Cobaea (Polemoniaceae). Syst. Bot. Monogr. 57: 1-81.
Prather, L. A., C. J. Ferguson, & R. K. Jansen. 2000. Polemoniaceae phylogeny and classification:
Implications of sequence data from the chloroplast gene ndhF. Amer. J. Bot. 87: 1300-1308.
Rzedowski, J. & R. McVaugh. 1966. La vegetacin de Nueva Galicia. Contr. Univ. Mich. Herb. 9(1):
1-123.
Rzedowski, J. & G. Caldern de Rzedowski. 1995. Familia Polemoniaceae. Flora del Bajo y de Regiones
Adyacentes 33: 1-41.
Standley, P. C. 1914. A revision of the genus Cobaea. Contr. U.S. Natl. Herb. 17: 448-458.
Vargas, A. & A. Prez. 1996. Estudios florsticos en Guerrero. No. 7. Cerro Chiletpetl y alrededores
(Cuenca del Balsas). Facultad de Ciencias, Universidad Nacional Autnoma de Mxico. Mxico,
D.F. 49 pp.
Vzquez, J. A., R. Cuevas, T. S. Cochrane, H. H. Iltis, F. J. Santana & L. Guzmn. 1995. Flora de
Manantln. Universidad de Guadalajara-Imecbio/University of Wisconsin-Madison. Sida,
Botanical Miscellany 13: 312.
Verduzco, C. & L. C. Rodrguez. 1995. Estudios florsticos en Guerrero. No. 4. El Rincn de la Va.
Facultad de Ciencias, Universidad Nacional Autnoma de Mxico. Mxico, D.F. 43 pp.
Weberbauer, A. 1945. El mundo vegetal de los Andes Peruanos. Ministerio de Agricultura, Per. Lima.
776 pp.

Recibido en julio de 1999.

Aceptado en noviembre de 2000.

37